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Research Article
Paraphlomis leigongshanensis (Lamiaceae), a new species from Guizhou, China
expand article infoWei-Hao Yao‡, Guo-Bin Jiang‡, De-Hui Yu§, Sheng Chen|, Hong-Mei Chen‡, Li Dai‡, Bai-Qiu He, He Li‡
‡ Key Laboratory of National Forestry and Grassland Administration on Biodiversity Conservation in Karst Mountainous Areas of Southwestern China, Guizhou Academy of Forestry, Guiyang, China
§ Guizhou Leigong Mountain National Nature Reserve Administration, Leishan, China
| Institute of Biology, Guizhou Academy of Sciences, Guiyang, China
¶ University of Chinese Academy of Sciences, Beijing, China

Corresponding author: Guo-Bin Jiang ( 1606580765@qq.com )

Academic editor: Chun-Lei Xiang
© 2025 Wei-Hao Yao, Guo-Bin Jiang, De-Hui Yu, Sheng Chen, Hong-Mei Chen, Li Dai, Bai-Qiu He, He Li.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Yao W-H, Jiang G-B, Yu D-H, Chen S, Chen H-M, Dai L, He B-Q, Li H (2025) Paraphlomis leigongshanensis (Lamiaceae), a new species from Guizhou, China. PhytoKeys 267: 23-35. https://doi.org/10.3897/phytokeys.267.157814
Open Access

Abstract

Paraphlomis leigongshanensis G.B. Jiang & W.H. Yao (Lamiaceae), a new species endemic to Leigong Mountain National Nature Reserve (Guizhou, China), is described and illustrated. Morphologically, it is most similar to P. jiangyongensis, but can easy be distinguished by its lamina shape, calyx teeth shape, and corolla color. A close relationship between the new species and P. gracilis var. lutienensis was revealed by molecular phylogenetic analyses based on ETS and ITS sequences, but they are morphologically distinct from each other. With only three known populations (ca. 200 mature individuals) in subtropical evergreen forests, the new species is assessed as Endangered (EN) under IUCN criteria B2ab(iii); D.

Key words:

Guizhou, Lamioideae, new taxon, phylogenetic analyses

Introduction

Paraphlomis (Prain) Prain (Lamiaceae, Lamioideae) is a member of the tribe Paraphlomideae Bendiksby, which was established to accommodate Paraphlomis, Matsumurella Makino, and Ajugoides Makino based on molecular phylogenetic evidence (Bendiksby et al. 2011; Zhao et al. 2021). Most species of Paraphlomis are distributed in southern China, with several taxa extending into the Himalayas, Korea, and Southeast Asia (Wu and Li 1977; Li and Hedge 1994a; Ko et al. 2014; Chen et al. 2021). Previous molecular phylogenetic studies revealed that Paraphlomis was not monophyletic, as species of Matsumurella were nested within it (Chen et al. 2021, 2022a; Guo et al. 2023). Recently, Yuan et al. (2024) treated Matsumurella and Ajugoides as synonyms of Paraphlomis based on plastome data and nuclear ribosomal DNA (nrDNA) sequences, as well as morphological evidence. Morphologically, Paraphlomis is characterized by its rhizomatous or stoloniferous habit, plants with simple hairs, nutlets with a truncate apex, and actinomorphic calyces (Wu and Li 1977; Bendiksby et al. 2011; Chen et al. 2021; Yuan et al. 2024). According to the Catalogue of Life China 2025 Annual Checklist and recent references, Paraphlomis comprises 51 species and six varieties (Li and Hedge 1994a; Chen et al. 2021; Yuan et al. 2024, 2025; Liu et al. 2025).

During a botanical field survey in Leigong Mountain National Nature Reserve, Guizhou Province, in 2024, we discovered an unknown species of Paraphlomis. This species is distinguished by its dwarf habit, a white corolla with pink stripes, and hairy mericarps. After careful field observations, morphological comparisons with other members of Paraphlomis, and molecular phylogenetic analyses, we confirmed its specific status and taxonomic placement within the genus. We hereby named it as P. leigongshanensis G.B. Jiang & W.H. Yao.

Materials and methods

Morphological study

During February 2024 and July 2025, four field surveys were carried out at the type locality of the new species. Specimens from several herbaria were examined, including GYBG, GZTM, HGAS, HIB, IBK, IBSC, KUN, NAS, and PE (herbarium abbreviations follow the conventions outlined by Thiers (2025)). Additionally, we consulted several online databases, including the China Plant Photography Database (http://ppbc.iplant.cn/), the Chinese Field Herbarium (http://www.cfh.ac.cn/), and Global Plants (http://plants.jstor.org/), to obtain supplementary information on species distributions and morphological data. To distinguish the new species from other species of Paraphlomis, we conducted detailed morphological comparisons, focusing on key characteristics such as stems, leaves, calyx, corolla, mericarps, and trichomes. All measurements were made using a dissecting microscope. The morphological description followed relevant taxonomic and floristic literature on Paraphlomis, with terminology standardized according to Li and Hedge (1994b), ensuring consistent and accurate morphological interpretation.

Molecular phylogenetic analyses

For the molecular phylogenetic reconstruction of Paraphlomis, we included a total of 53 samples: 38 representing Paraphlomis taxa (34 species and four varieties) and two outgroup taxa. The outgroups comprised Phlomoides dentosa var. glabrescens (Danguy) C.L. Xiang & H. Peng and Phlomis fruticosa L. Only one accession of the new species was sampled and sequenced in this study, while sequence data for other species were obtained from GenBank and previous studies (Chen et al. 2021, 2022a, 2022b).

Genomic DNA was extracted from dried leaf tissue using a modified CTAB method (Doyle and Doyle 1987). Previous studies have shown that phylogenetic trees based on nuclear ribosomal DNA internal and external transcribed spacers (nrITS and nrETS) offer more effective insights into the phylogenetic relationships of Paraphlomis than those constructed using chloroplast DNA (Chen et al. 2021; Yuan et al. 2024). Thus, only the nrDNA data was used here for phylogenetic reconstruction. Voucher information and GenBank accession numbers for all sequences are provided in Appendix 1.

The best-fitting substitution model was selected using the Akaike Information Criterion (AIC) based on sequence data in jModelTest v.2.1.7 (Darriba et al. 2012). Phylogenetic analyses were performed using Bayesian Inference (BI) and Maximum Likelihood (ML) methods, employing MrBayes (Ronquist et al. 2012) and RAxML v.8.2.9 (Stamatakis 2014) on the CIPRES Science Gateway v.3.3 (Miller et al. 2010), respectively. The Markov chain Monte Carlo algorithm was run for 20 million generations with four incrementally heated chains starting from random trees and sampling one out of every 1,000 generations. The initial 25% of sampled trees were discarded as burn-in and the 50% majority-rule consensus tree was calculated from remaining trees with nodal support summarized as posterior probabilities (PP). All phylogenetic trees were visualized and edited for clarity using FigTree v.1.4.2 (Rambaut 2014), with PP and bootstrap support (BS) values annotated on the branches.

Results and discussion

Morphological comparison

Morphological comparisons were based on an examination of 196 specimens representing 14 species of Paraphlomis. A key diagnostic character of the potential new species P. leigongshanensis is the hairy mericarps, which serves as an important taxonomic feature for the infrageneric classification (Chen et al. 2021; Yuan et al. 2024). Morphologically, P. leigongshanensis is most similar to P. jiangyongensis in the dwarf habit ascending stems, and hairy mericarps (Table 1). However, it differs from P. jiangyongensis by its abaxially purple (vs. green) laminae, calyx teeth acuminate (vs. acute) at apex, and longer corolla tubes. Additionally, the corolla of P. leigongshanensis is white with pink stripes and spots, with the markings concentrated on the lower lip, differing from the pale yellow corolla lips with red spots in P. jiangyongensis. This species is distributed in the mid-subtropical evergreen broad-leaved forests in Leishan County, Guizhou, exhibiting geographical isolation from P. jiangyongensis (Appendix 2: Fig. A1).

Table 1.
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Morphological comparisons among P. leigongshanensis, P. jiangyongensis, P. gracilis var. gracilis, and P. gracilis var. lutienensis.

Character P. leigongshanensis P. jiangyongensis P. gracilis var. gracilis P. gracilis var. lutienensis
Stem Prostrate to ascending, 5–10 cm tall Ascending, 5–30 cm tall Erect, 30–100 cm tall Erect, 30–100 cm tall
Lamina Ovate to long elliptic, 2.5–7 × 2–5 cm, abaxially purple, densely villose Ovate to subcircular, 3–8 × 2–5 cm, abaxially light green, densely strigose Lanceolate, 5–10 × 1.7–3.3 cm, abaxially green, densely strigose and glandular Lanceolate, 6.5–10.5 × 1.7–2.5 cm, abaxially green, densely strigose and glandular, subsessile and narrowly lanceolate
Verticillaster 2–10-flowered 2–8-flowered (2–) 4–8 (–12)-flowered (2–) 4–8 (–12)-flowered
Pedicel Sessile Sessile Subsessile Ca. 4 mm long
Calyx Purplish–red, densely villose Green, densely strigose Green, densely retrorse strigose Green, densely retrorse strigose
Calyx teeth 4–8 mm long, triangular, apex acuminate Ca. 2 mm long, triangular, apex acute Ca. 6 mm long, triangular, apex acuminate or subulate Ca. 4 mm long, triangular, apex acuminate
Corolla 12–17 mm long, white lower lip with pink stripes and spots 7.5–10 mm long, pale yellow, lower lip with red spots 15 mm long, upper yellow, white or yellowish green, lower lip white with purple spots or yellow with red spots 15 mm long, yellow, lower lip with pink spots

Molecular phylogenetic analyses

The concatenated nrDNA dataset had a total length of 1259 bp, comprising 814 bp from the ITS region and 445 bp from the ETS region. The dataset contained 356 variable sites (168 from ITS, 188 from ETS) and 205 parsimony-informative characters (89 from ITS, 116 from ETS).

The resulting phylogenetic tree of Paraphlomis was consistent with previous studies (Chen et al. 2021; Yuan et al. 2024). The potential new species P. leigongshanensis was resolved as sister to P. gracilis var. lutienensis (Y. Z. Sun) C. Y. Wu (Fig. 1: BS = 100%/PP = 1.00), and the P. leigongshanensis–P. gracilis var. lutienensis clade was further sister to P. gracilis var. gracilis Kudô (Fig. 1: BS = 100% / PP = 1.00). The three taxa were all nested within the “Clade III” proposed by Chen et al. (2021) and Yuan et al. (2024) (Fig. 1: BS = 83%/PP = 1.00). Species in this clade (highlighted in blue in Fig. 1) are all characterized by hairy mericarps (Chen et al. 2021; Yuan et al. 2024). Despite the close relationships and geographical affinities (Appendix 2: Fig. A1) among P. leigongshanensis, P. gracilis var. gracilis, and P. gracilis var. lutienensis, P. leigongshanensis is morphologically distinct from the latter two taxa. Specifically, P. leigongshanensis exhibits a creeping habit with stems 5–10 cm tall, ovate to elliptic leaves, reddish-purple calyces, sessile flowers, and white corollas with pink stripes and spots. In contrast, P. gracilis var. gracilis has erect and much taller stems (up to 1 m tall), lanceolate leaves, green calyces, subsessile flowers, and yellowish green to white corollas with purple spots (Table 1). Paraphlomis gracilis var. lutienensis differs from both species by its subsessile and narrowly lanceolate laminae and yellow corollas. Moreover, a recent phylogenetic study based on plastome data (Yuan et al. 2024) indicated that the phylogenetic relationship between P. gracilis var. gracilis and P. gracilis var. lutienensis is consistent with the results of our study. Based on our phylogenetic results, although P. leigongshanensis is grouped with P. gracilis var. gracilis and P. gracilis var. lutienensis, there is sufficient evidence to support its recognition as a distinct species. The morphological differences further reinforce its taxonomic distinctiveness and provide additional support for its designation as a new species.

Figure 1. 

Bayesian 50% majority-rule consensus tree of Paraphlomis based on combined nuclear (ITS and ETS) data set. Support values ≥ 0.50 PP or 50% BS are displayed above the branches (an “*” indicates a support value of 1.00 PP or 100% BS). Multiple accessions of the same species are numbered according to Appendix 1

Taxonomic treatment

Paraphlomis leigongshanensis G.B.Jiang & W.H.Yao, sp. nov.

urn:lsid:ipni.org:names:77365161-1
Figs 2, 3; Appendix 2: Figs A1, A2 雷公山假糙苏

Type.

China • Guizhou Province: Qiandongnan Miao and Dong Autonomous Prefecture, Leishan County, Leigong Mountain National Nature Reserve, 26°24'30.91"N, 108°15'53.77"E, alt. 1104 m, 23 Aug. 2024, G.B. Jiang, S. Chen, H.M. Chen & D.H. Yu GB2024156 (holotype: GF 09040011!; isotypes: KUN 1644009!, HGAS 0128505!).

Diagnosis.

Paraphlomis leigongshanensis is morphologically most similar to P. jiangyongensis but can be distinguished by its calyx purplish red (vs. green), apex of calyx teeth lanceolate (vs. acute), corolla white with pink stripes (vs. pale yellow with red spots), and corolla tube 12–17 mm long (vs. 7.5–10 mm long).

Figure 2. 

Paraphlomis leigongshanensis from the type locality. A. Habitat; B-D. Habit; E. Inflorescence; F. Flowering condition; G. Adaxial view of leaf; H. Abaxial view of leaf; I. Stem; J. Calyces; K. Dissected corolla and pistil; L. Corollas; M. Dissected calyx and mericarps; N. Dried mericarps. (Photographed by Guo-Bin Jiang).

Description.

Perennial herb, 5–10 cm tall, stoloniferous. Stems simple, prostrate to ascending, slightly 4-angled, densely villose. Leaves opposite, ovate to elliptical, papery, 2.5–8 × 2–5 cm; margin shallowly crenate, apex obtuse to subrounded, base broadly cuneate to rounded; adaxial surface green, abaxial purple, densely villose on both surfaces, particularly along veins, lateral veins 3–5-paired; petioles 0.5–3.5 cm long, densely villose. Verticillasters 2–10-flowered, sessile; bracteoles minute, early deciduous. Calyx purplish red, tubular-campanulate, 10–15 mm long, densely villose outside, glabrous inside; teeth 5, subequal, triangular, erect, 4–8 mm long, apex acuminate. Corolla white with pink stripes and spots, 2–3 cm long, densely pubescent outside; tube 1.2–1.7 cm long, slightly dilated at throat, pubescent annulate inside at 1/3 distance from base; bilabiate, upper lip pink, oblong, entire, erect, concave, 8–10 mm long, ca. 5 mm wide, lower lip pink, 3-lobed, medium lobe subcircular with pink stripes and spots, apex slightly emarginate, 6–7 mm wide, lateral lobes ovate, ca. 3.5 mm wide. Stamens 4, straight, included; filaments flat, pubescent at base; anther cells 2, parallel. Style included, glabrous, apex subequally 2-lobed, lobes subulate. Ovary densely pubescent and glandular at apex. Mericarps 4, dark brown, triquetrous-oblong, ca. 3.6 mm long, densely pubescent at apex.

Figure 3. 

Line drawing of Paraphlomis leigongshanensis. A. Plant; B. Flower; C. Dissected corolla and pistil; D. Calyx; E. Dissected calyx; F. Nutlets. (Drawn by Hong-Mei Chen).

Distribution, habitat and conservation status.

Paraphlomis leigongshanensis is currently known from only three populations in adjacent valleys approximately 100 m apart, within Leigong Mountain National Nature Reserve, Guizhou, China (Appendix 2: Fig. A1). The total population comprises about 200 mature individuals. The species occurs in the subtropical monsoon climate zone, with populations confined to the area near Getou Village, and is vulnerable to anthropogenic disturbances. Based on the IUCN Red List criteria B2(a, b(iii)) (extent of occurrence < 500 km2; < 5 locations; continuing habitat degradation), D (Estimated population size < 250 mature individuals), we proposed an Endangered (EN) assessment (IUCN 2024). Since similar adjacent habitats may harbor undiscovered populations, available evidence did not support Critically Endangered (CR) status.

Phenology.

Flowering was observed from May to August, and fruiting from August to October.

Etymology.

The specific epithet “leigongshanensis” is derived from the type locality of the new species, i.e., Leigong Mountain National Nature Reserve, Leishan County, Guizhou Province, China.

Chinese name (assigned here).

léi gōng shān jiă cāo sū (雷公山假糙苏). The name means a species of Paraphlomis only found in Leigong Mountain National Nature Reserve, Guizhou Province, China.

Additional specimens examined (paratypes).

China • Guizhou Province: Qiandongnan Miao and Dong Autonomous Prefecture, Leishan County, Leigong Mountain National Nature Reserve, 26°24'19.92"N, 108°15'19.20"E, alt. 1100 m, 1 Jul. 2025, S. Chen & L. Hao GB2025015 (GF 09040012!, KUN 1644008!).

Acknowledgements

We are grateful to Dr. Ya-Ping Chen of the Kunming Institute of Botany, Chinese Academy of Sciences, for conducting the phylogenetic analysis. We thank Tao-Hua Yuan (PhD candidate, Tibet University), Xiao-Kai Xiong (Master’s candidate, Guizhou University), and Jiang-Miao Gu (Master’s candidate, Guizhou University) for their suggestions on the manuscript.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of AI

No use of AI was reported.

Funding

This study was financially supported by the Survey and Assessment of Newly Added National Key Protected Wild Plant Resources in Guizhou Province (Three stage) (MCHC-ZD20242057), Guizhou Provincial Basic Research Program (Qiankehe Fundamentals-ZK [2023] General 234), and Guizhou Provincial Basic Research Program (Qiankehe Fundamentals-MS [2025] 327).

Author contributions

Guo-Bin Jiang conceptualized and led the project. Field investigation and sample collection were conducted by Guo-Bin Jiang, Sheng Chen, De-Hui Yu, Hong-Mei Chen, Li Dai, and Wei-Hao Yao. Wei-Hao Yao, Guo-Bin Jiang, Hong-Mei Chen, and Bai-Qiu He examined and detailed the fine structure of the species represented in the study. The initial draft of the manuscript was prepared by Wei-Hao Yao and Guo-Bin Jiang. All authors approved the final manuscript.

Author ORCIDs

Wei-Hao Yao https://orcid.org/0009-0001-8231-0867

Guo-Bin Jiang https://orcid.org/0000-0002-4631-7906

De-Hui Yu https://orcid.org/0009-0008-4264-8995

Sheng Chen https://orcid.org/0000-0002-2727-6504

Hong-Mei Chen https://orcid.org/0009-0002-2368-7813

Li Dai https://orcid.org/0009-0004-3907-8582

Bai-Qiu He https://orcid.org/0009-0009-7491-3586

He Li https://orcid.org/0009-0006-0597-3209

Data availability

The newly-obtained ITS and ETS sequences have been submitted to the NCBI (accession numbers: PV476761; PV478008). All of the other data that support the findings of this study are available in the main text.

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Appendix 1

Table A1.
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Sequence information for all samples used in present study. A “/” indicates a missing sequence. Herbarium abbreviations are listed after the vouchers. The accession numbers marked in bold represent sequences newly generated.

Taxon Voucher Country ITS ETS
Paraphlomis albida Hand.-Mazz. var. albida A. Liu et al. LK0841 (CSFI) Ningyuan, Hunan, China MW602124 MW602091
Paraphlomis albida var. brevidens Hand.-Mazz. Y.P. Chen EM312 (KUN) Hezhou, Guangxi, China MW602130 MW602098
Paraphlomis albiflora (Hemsl.) Hand.-Mazz. C.M. Tan et al. 1806393 (JJF) Jiujiang, Jiangxi, China / MW602101
Paraphlomis baiwanensis W.Y. Zhao, Y.P. Chen & Q. Fan 1 Y.S. Chen et al. QY20230302 (IBSC) Qingyuan, Guangdong, China PP897029 PP897950
Paraphlomis baiwanensis W.Y. Zhao, Y.P. Chen & Q. Fan 2 Q. Fan et al. 20251 (SYS) Qingyuan, Guangdong, China PP897030 PP897951
Paraphlomis caloneura K.J. Yan, Y.P. Chen & Y.F. Huang W.H. Wu et al. LHT1841 (KUN) Napo, Guangxi, China OQ627454 OQ628080
Paraphlomis chinensis (Benth.) J.C. Yuan, Y.P. Chen & C.L. Xiang 1 Y. Yang OYY00316 (KUN) Pingxiang, Jiangxi, China MW602147 MW602117
Paraphlomis chinensis (Benth.) J.C. Yuan, Y.P. Chen & C.L. Xiang 2 Y. Yang OYY00131 (KUN) Guilin, Guangxi, China MW602148 MW602118
Paraphlomis coronata (Vaniot) Y.P. Chen & C.L. Xiang 1 E.D. Liu et al. 3043 (KUN) Emeishan, Sichuan, China MW602137 MW602107
Paraphlomis coronata (Vaniot) Y.P. Chen & C.L. Xiang 2 C.L. Xiang 358 (KUN) Jiangkou, Guizhou, China MW602123 MW602090
Paraphlomis foliata (Dunn) C.Y. Wu & H.W. Li S.P. Chen s.n. (KUN) Jiangle, Fujian, China / MW602097
Paraphlomis gracilis (Hemsl.) Kudô var. gracilis 1 A. Liu LK0931 (CSFI) Changsha, Hunan, China MW602134 MW602104
Paraphlomis gracilis (Hemsl.) Kudô var. gracilis 2 C.L. Xiang XCL1315 (KUN) Chongqing, China MW602141 MW602111
Paraphlomis gracilis var. lutienensis (Y.Z. Sun) C.Y. Wu C.L. Xiang XCL881 (KUN) Shibing, Guizhou, China MW602131 MW602099
Paraphlomis hirsutissima C.Y. Wu & H.W. Li F. Zhao & G. Chen XCL2115 (KUN) Malipo, Yunnan, China OQ627453 OQ628079
Paraphlomis hispida C.Y. Wu X. Li LX200702 (GXF) Napo, Guangxi, China MW602132 MW602102
Paraphlomis hsiwenii Y.P. Chen & Xiong Li 1 W.H. Wu et al. DD426 (KUN) Jingxi, Guangxi, China OP605346 OP609841
Paraphlomis hsiwenii Y.P. Chen & Xiong Li 2 W.H. Wu et al. DD426 (KUN) Jingxi, Guangxi, China OP605347 OP609842
Paraphlomis intermedia C.Y. Wu & H.W. Li X. Zhong et al. ZX16823 (CSH) Suichang, Zhejiang, China MW602135 MW602105
Paraphlomis javanica (Blume) Prain var. javanica 1 Y.P. Chen s.n. (KUN) Kunming, Yunnan, China MW602121 MW602088
Paraphlomis javanica (Blume) Prain var. javanica 2 L.B. Jia et al. JLB0029 (KUN) Maguan, Yunnan, China MW602143 MW602113
Paraphlomis javanica var. pteropoda D. Fang & K.J. Yan X. Li 2020090501 (GXF) Jingxi, Guangxi, China MW602140 MW602110
Paraphlomis jiangyongensis X.L. Yu & A. Liu 1 A. Liu et al. LK1104 (CSFI) Jiangyong, Hunan, China MW602128 MW602095
Paraphlomis jiangyongensis X.L. Yu & A. Liu 2 A. Liu et al. LK1104 (CSFI) Jiangyong, Hunan, China MW602129 MW602096
Paraphlomis kwangtungensis C.Y. Wu & H.W. Li Q. Fan et al. 19738 (SYS) Qujiang, Guangdong, China PP713070 PP706067
Paraphlomis lanceolata Hand.-Mazz. 1 C.Z. Huang s.n. (KUN) Guidong, Hunan, China MW602145 MW602115
Paraphlomis lanceolata Hand.-Mazz. 2 A. Liu et al. LK0825 (CSFI) Ningyuan, Hunan, China MW602146 MW602116
Paraphlomis lancidentata Y.Z. Sun X. Zhong et al. ZX16824 (CSH) Suichang, Zhejiang, China MW602136 MW602106
Paraphlomis leigongshanensis G.B. Jiang & W.H. Yao G.B. Jiang et al. GB2024156 (GF) Leishan, Guizhou, China PV476761 PV478008
Paraphlomis longicalyx Y.P. Chen & C.L. Xiang Y.P. Chen et al. EM583 (KUN) Huanjiang, Guangxi, China OK104771 OK104774
Paraphlomis membranacea C.Y. Wu & H.W. Li M.S. Nuraliev 1057 (MW) Thanh Son, Phu Tho, Vietnam / MW602100
Paraphlomis montigena (X.H. Guo & S.B. Zhou) J.C. Yuan, Y.P. Chen & C.L. Xiang Y.C. Dai s.n. (KUN) Hangzhou, Zhejiang, China OM836064 OM884453
Paraphlomis nana Y.P. Chen, C. Xiong & C.L. Xiang 1 C. Xiong XC21097 (KUN) Chengkou, Chongqing, China OM836062 OM884451
Paraphlomis nana Y.P. Chen, C. Xiong & C.L. Xiang 2 C. Xiong & H.L. Zhou XC21126 (KUN) Wushan, Chongqing, China OM836063 OM884452
Paraphlomis octopus Q. Fan, Y.P. Chen & Ying Liu 1 Y.P. Chen & Y. Zhao EM1391 (KUN) Huaiji, Guangdong, China MW602126 MW602093
Paraphlomis octopus Q. Fan, Y.P. Chen & Ying Liu 2 Q. Fan et al. 19752 (SYS) Fengkai, Guangdong, China PP713071 PP706068
Paraphlomis octopus Q. Fan, Y.P. Chen & Ying Liu 3 Q. Fan et al. 19760 (SYS) Pingle, Guangxi, China PP713072 PP706069
Paraphlomis pagantha Dunn 1 L.X. Yuan et al. s.n. (KUN) Qionghai, Hainan, China OP605345 OP609840
Paraphlomis pagantha Dunn 2 X.Y. Jiang et al. HN001 (SYS) Wenchang, Hainan, China PP713073 PP706070
Paraphlomis patentisetulosa C.Y. Wu C.L. Su et al. XY015 (KUN) Xinyi, Guangdong, China OQ627455 OQ628081
Paraphlomis paucisetosa C.Y. Wu 1 X.X. Zhu s.n. (KUN) Malipo, Yunnan, China MW602125 MW602092
Paraphlomis paucisetosa C.Y. Wu 2 X. Li LX200704 (GXF) Napo, Guangxi, China MW602133 MW602103
Paraphlomis qingyuanensis W.Y. Zhao, R.M. Wu & Q. Fan 1 Q. Fan et al. QYK-HH-1904 (SYS) Yingde, Guangdong, China PP897031 PP897952
Paraphlomis qingyuanensis W.Y. Zhao, R.M. Wu & Q. Fan 2 Q. Fan et al. 20255 (SYS) Yingde, Guangdong, China PP897032 PP897953
Paraphlomis reflexa C.Y. Wu & H.W. Li Z.Z. Yang et al. s.n. (HIB) Tongshan, Hubei, China MW602122 MW602089
Paraphlomis strictiflora J.C. Yuan, B. Chen & C.L. Xiang B. Chen et al. CB05956 (CSH) Yinjiang, Guizhou, China / OP609839
Paraphlomis subcoriacea C. Y. Wu ex H. W. Li 1 Q. Fan et al. QYK-CJ-1911 (SYS) Yangshan, Guangdong, China PP897033 PP897954
Paraphlomis subcoriacea C. Y. Wu ex H. W. Li 2 Q. Fan et al. QYK-LB-1920 (SYS) Yangshan, Guangdong, China PP897034 PP897955
Paraphlomis yangsoensis (Y.Z. Sun) J.C. Yuan, Y.P. Chen & C.L. Xiang L. Wu & W.B. Xu 10965 (IBK) Yangshuo, Guangxi, China MW602142 MW602112
Paraphlomis yingdeensis W.Y. Zhao, Y.Q. Li & Q. Fan Q. Fan et al. 19013 (SYS) Yingde, Guangdong, China OP605348 OP609843
Paraphlomis youyangensis H. Jiang, R.B. Zhang & Tan Deng R.B. Zhang et al. ZRB2531 (ZY) Youyang, Guizhou, China OR488121 OR487461
Phlomis fruticosa L. Y. Tong s.n. (KUN) Shanghai, China (cultivated) MW602119 MW602086
Phlomoides dentosa var. glabrescens (Danguy) C.L. Xiang & H. Peng Y.P. Chen EM360 (KUN) Beijing, China (cultivated) MW602120 MW602087

Appendix 2

Supplementary figures provided in the present study.

Figure A1. 

Distribution of Paraphlomis leigongshanensis, P. jiangyongensis, P. gracilis var. gracilis, and P. gracilis var. lutienensis.

Figure A2. 

Holotype (A) and isotype (B) specimens of Paraphlomis leigongshanensis.

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